The most well-known
late archaic humans were the Neandertals.
More Neandertal skeletons have been found than any other ancient human
species. They lived in Europe and Southwest Asia from
at least 130,000 years ago until
around 28,000 years ago.
This is in the late Pleistocene Epoch.
It is likely that the Neandertals evolved from Homo heidelbergensis in Southern Europe. Neandertal-like
skull characteristics have been found in 400,000 year old fossils from Spain.
The Neandertals adapted physically and culturally to the ice age conditions
that prevailed during much of their time.

No other ancient people have aroused more controversy and
confusion over the last century and a half than have the Neandertals. There is an on-going debate as to whether they should be consideredHomo
sapiens. If they were members of our species, they were a different
variety or race (Homo sapiens neanderthalensis). On the other hand, if
they were dissimilar enough to be a distinct
species, they should be called Homo neanderthalensis.

Important Neandertal
Sites

Date of Fossil(years
ago)

Croatia:

Krapina

130,000-120,000

Vindija

44,400-38,300

France:

La Ferassie

75,000-60,000

La Chapelle-aux-Saints

56,000 or 47,000

La Quina

75,000-40,000

Le Moustier

40,000

St Césaire

36,000

Germany:

Neandertal

40,000

Gibraltar:

Forbe's Quarry

70,000-45,000?

Gorham's Cave

32,000-24,000

Italy:

Saccopastore

129,000-122,000?

Mt. Circeo

60,000-55,000

Date of Fossil(years
ago)

Iraq:

Shanidar

65,000-35,000

Israel:

Tabun

103,000-80,000

Kebara

60,000

Amud

55,000-47,000

Romania:

Peştera Muierii

32,000

Peştera cu Oase

30,000

Russia:

Mezmalskaya

70,000-60,000

Okladnikov

38,000-30,000

Spain:

El Sidrón

43,000

Uzbekistan:

Obi-Rakhmat
and Anghilak

100,000-53,000?

Neandertal
Discoveries

The first
discovery of Neandertal bones may have been during the late 1820's in Western
Europe. Subsequently, other, better documented Neandertals were
found and stored in museums without being recognized as early forms
of humans. In 1839, for instance, a portion of a child's skeleton was
found in Engis, Belgium. In 1848, an adult female skull was found at
Gibraltar, at the southern tip of Spain.

Johan Karl Fuhlrott
1803-1877

In 1856 a discovery
was made in Germany that finally sparked the recognition that these were, in
fact, not just strange looking modern people. This was
the discovery of a skull and a number of other bones from a limestone cave
deposit in the small Neander River Valley near Dïüsseldorf. Thinking that they
were from a bear, the quarrymen gave them to a local school teacher and amateur naturalist,
Johan
Karl Fuhlrott. He recognized them as being human but somewhat different
from those of modern Europeans. When several leading paleontologists
and medical pathologists in Germany became aware of the fossils, a disagreement
developed about who the "Neandertal Man" might have been.
It was suggested that he had been an old Roman, a Dutchman, and even a Central
Asian soldier in the service of the Russian czar during the Napoleonic
wars of the
early 19th century. The reality that these bones were from an earlier
variety or species of human was not yet conceivable to most
of the scientific world in
the 1850's.

What finally
convinced the scientific community that Neandertals were very ancient
Europeans was a combination of additional fossil discoveries and
new perspectives that largely began with the publication of Charles
Darwin's On the Origin of Species in 1859. This seminal work in
biology popularized the idea that species of living things evolve over time
as a result of natural selection. Subsequently, it was not a major
leap in understanding to realize that humans also must have evolved from
earlier forms. In fact, Darwin proposed just that in his 1871
influential publication The Descent of Man, and Selection in
Relation to Sex. For enlightened
Victorians, the Neandertals ultimately came to be seen as important human
ancestors. They were given the name "Neandertal Man" in 1864
by William King, an Irish anatomist. He named them after the Neander
River Valley (or Tal, in German).

NOTE:
In many older books, Neandertal is spelled with a "thal" ending
(Neanderthal) instead of "tal".
This is the Old German spelling that was replaced in the early 20th century.
However, the outdated usage of
"thal" persists in some English publications.
It is also continued in scientific classification (Homo neanderthalensis)
because it was used when Neandertal's were first described in the 19th
century. In this tutorial series, the modern
"tal" is used.

In 1886, two Neandertal skeletons were found
in a Belgium cave site named Spy. These specimens were physically associated
with stone tools and food refuse bones from extinct
ice age animals. This evidence once and for all demonstrated that
Neandertals were not modern people. As a result, the
majority of European paleontologists in the late 1880's finally accepted them for what they really were.

Marcellin Boule
1861-1942

The remains of more
than 400 Neandertals have been found. The most controversial one was
excavated in 1908 at La Chapelle-aux-Saints
in southwestern France. This is a nearly
complete skeleton of a man who would have been elderly by Neandertal
standards. The bones were analyzed between 1911 and 1913 by the noted
French paleontologist, Marcellin Boule . Unfortunately, Boule's
prejudices got in the way of scientific objectivity. He described the La
Chapelle-aux-Saints man, and subsequently all Neandertals, as dull-witted,
brutish, ape-like creatures who walked hunched over with a shuffling gait.
Unfortunately, this mistaken view was universally accepted by paleoanthropologists for decades. It also
became the source of the popular images of dim-witted cavemen that still
appear in cartoons and movies.

Diorama of Neandertals
in an American
museum
during the 1930's reflecting
the misconception
reinforced
by Marcellin Boule's description
of
them as dull-witted, brutish,
ape-like creatures.

After reanalysis of the La Chapelle-aux-Saints
skeleton in the 1950's, it became clear that a serious mistake had been
made. This had been an atypical Neandertal. He was at least
40 years old with a somewhat hunched over posture resulting from severe arthritis
in his spine. There was a bowing of his legs that may have resulted from
rickets disease
in childhood. He had lost most of his teeth and
part of his jaw resulting in a disharmonic looking face. Despite these
deforming infirmities, it is now clear that the La Chapelle-aux-Saints man was
much more like us in appearance, intelligence, and physical
ability than had been believed by
Marcellin Boule.

La Chapelle-aux-Saints
man

Analysis
of Neandertal Anatomy

The Neandertals were physically diverse, but
in general they were larger boned
and more heavily muscled than most modern humans. This was particularly
true of the European Neandertals, like the La
Chapelle-aux-Saints man. Some
of the Southwest Asian Neandertals were less robust in appearance
and somewhat more like modern humans. The Neandertals were relatively short
and stocky compared to some other archaic humansand modern
Europeans. Adult male Neandertals averaged 5 feet 5
inches
tall (164 cm.) and 143 pounds (65 kg). Females
averaged 5 feet 1 inch tall (155 cm) and 119 pounds (54 kg). They probably stood as erect as we do and were
fully bipedal. They were not
only strong but apparently quite flexible. The thickness and high density
of their
leg bones suggest that they did a great deal of walking and running.
Their lower arm and leg bones were short compared to modern humans.
These traits were likely adaptations to an aggressive hunting and gathering
way of life as well as to the cold climates in which most Neandertals lived.
The fact that adult Neandertal skeletons frequently have multiple healed bone
fractures suggests that these people had rough lives. Some researchers
believe that many of the broken bones were the result of hunting large game
animals up close with jabbing spears--a dangerous enterprise.

Reconstruction of the
Neandertal appearance

It is likely that at least some Neandertals had pale skin
color and red hair. This is based on the discovery of a variant of the
MC1R gene associated with these traits in the bones of two European
Neandertals dated to around 50,000 years ago. This was very likely an
adaptation that helped their bodies produce more Vitamin D and subsequently
absorb more calcium from their food in ice age Europe.

Neandertal heads were long (from front to back) compared to ours. This
resulted in relatively low, sloping foreheads. At the back of
their skulls, they had a prominent bulge or projection called an occipital bun. They had
large faces (especially in the middle part) with big noses andprominent brow
ridges that extended between the eyes. They lacked the pointed chin that is common in modern
Homo sapiens.
These traits give the Neandertal face and head an appearance more reminiscent
of late Homo erectusand Homo heidelbergensisthan of modern people.

Neandertal

Modern human

The brain size of Neandertals was close to that of modern
humans, and the structural organization of their brains was essentially the
same as well. The average Neandertal brain was actually somewhat
larger than the brains of most people today. However, the difference is
minimal
when people of similar body size are compared. In fact, the average
Neandertal brain may have been slightly smaller from this perspective. The
large heads and
stocky bodies of Neandertals very likelywere more efficient in cold climatesand
were probably selected for
by nature.This trend has been observed among contemporary
Native American populations
living in sub-arctic environments. A larger head and
more compact body shape potentially produce more body heat relative to the
amount that is lost to the environment through radiation. A
bigger brain carries a high energy overhead. The human brain uses
around 20% of the energy that we get from our food when we are resting, but
it is only 2% of our body mass. This is one reason that the brains of
new-born humans are only about 25% the size of those of adults. It is
hard for a pregnant woman's body to feed her own brain and that of her baby
at the same time. It is even more difficult when
there are
twins.

Comparison
of Cranial Capacities

range (cm3)

average
(cm3)

chimpanzees

300-500

----

australopithecines

400-515

----

Homo habilis

509-752

612

Homo erectus

750-1250

930

Homo heidelbergensis

1100-1390

1206

Neandertals

1200-1750

1450

modern Homo sapiens

900-1880

1345

Note:There is a considerable range in body
and head size
among modern Homo sapiens around the world. As a
consequence, the average brain size is smaller than would
initially seem likely. However, the average for some modern
populations (especially European and most African ones)
is slightly larger than that of Neandertals.

Female Neandertal brains were about 200
cm3
smallerthan those of males.
This sexual dimorphism should not be a surprise since female bodies were
smaller. Modern human female brains are about 10% smaller than those
of males for the same reason.

NOTE:
It would be a mistake to assume that a minor difference in overall brain size
is directly correlated
with intelligence among archaic or modern
humans. However, the gross difference in
cranial capacity between the earliest human species 2.5
million years ago and recent Homo sapiens
probably does reflect potential intelligence differences. In order to
trace the development of intelligence, speech, and other mental capabilities,
it is more useful to examine changes in specific brain regions
and the genes that control their development.

It is now clear that upright
bodies and bipedal locomotion long preceded the evolution of the large human
brain. The early 20th century speculation that our
ancestors would be large brained apes proved to be incorrect. We
attained the full human form of bipedalism by about 2.5 million years
ago,if not earlier. However,
the size of our
brains continued to increase in a punctuated
evolutionary pattern. There apparently was a period of comparative stasis
beginning around 1.8 million years ago. However, by 800,000-600,000 years
ago, human brain size began to grow very
rapidly. This skyrocketing trend continued until
around 100,000years ago or a bit
earlier.

In other words, there was a
mosaic
pattern of evolution. We continued to evolve above the neck after
the rest of the body essentially reached its modern form. This process
of the brain increasing in size over and beyond that explainable by an
increase in body size has been referred to as encephalization.
The overall increase in
brain size was, in fact, mostly a result of changes in particular regions of
the cerebrum,
where most high level brain functions occur. It
is likely that nature was selecting
for the mental capabilities needed to adapt rapidly to new
environments. The
brain was being neurally reorganized for processing complex information. This can be seen indirectly in the evolution of
culture. Progressively more sophisticated stone tools are, in effect, the
material products of encephalization.

Who Were the Neandertalsand What Happened to Them?

We are still left with the question of whether
Neandertals were members of our species or another species with whom we share
a distant common ancestor. Two sources of evidence have shed light on
this issue--DNA and bones.

In 2009, a first draft of the Neandertal genome was completed.
It consists of 60% of their approximately 3 billion DNA base units.
They were sequenced mostly from bones found in Vindija Cave in Croatia.
Based on this information, Svante Paabo of the Max Planck Institute for
Evolutionary Anthropology in Germany concluded that the Neandertal and
modern human genomes share 99.5-99.9% of their base unitsequences. He estimated that the Neandertal line
began to diverge from ours by about 800,000 years ago and that we were
"genetically distinct" by 300,000 years ago. Further
analysis of the Vindija Cave Neandertal DNA by Richard Green of the
University of California, Santa Cruz led him to announce in 2010 that 1-4%
of the DNA in modern Europeans and Asians came from Neandertals.
Therefore, he suggested that there must have been some
interbreeding between modern humans and Neandertals around 87-37,000 years
ago. This most likely occurred in Southwest Asia, shortly after modern
humans migrated out of Africa. This would account for the lack of
Neandertal DNA markers in Sub-Saharan African populations today. The
fact that many North Africans also share Neandertal DNA is likely the result
of back migrations from Southwest Asia.

The
Neandertal In Us--the
difficulty of finding and extracting fossil DNA
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(length =
7 mins,
37 secs)

The Neandertal DNA in modern human
populations includes some of the genes for our HLA (human leukocyte antigen) immune system. It
has been suggested that this gave early modern human immigrants to Europe
and Asia critical protection to diseases that had not existed in their
African homeland. Mating with Neandertals and other archaic humans in
Eurasia would have helped the modern Homo sapiens adapt to viruses, bacteria,
and parasitic worms that they had not been exposed to before.

Supporting evidence of
interbreeding comes from 4 Neandertal skulls found in different
locations in Europe. They appear to have a mixture of Neandertal and modern
Homo sapiens anatomical characteristics. The implication is
that some of the Neandertals interbred with modern humans
resulting in gene flow between the
populations. If
that is true, then the genetic difference between us and them must not have
been as great as would be expected between two distinct species. In
other words, this would suggest that the Neandertals were a
distant variety of Homo
sapiens rather than a distinct species. One of the presumed Neandertal/modern human hybrids was a 4 year old child found in Portugal in
1999. What is striking about this fossil is that it dated to 24,500
years ago--3-4,000
years after the last known Neandertal.

The size of the total Neandertal
population was never very large and appears to have diminished steadily beginning around 35,000 years
ago. It is probable that at its maximum their widely dispersed
population had only around 70,000 individuals with about 7,000 breeders. The last secure date for a Neandertal site was about 28,000 years
ago. What happened to them is not clear. However, their
relatively abrupt disappearance roughly coincides with the arrival and
rapid growth in numbers of modern humans in Europe. A common view is that Neandertals could not compete effectively with the
technologically more advanced and numerically larger new-comer population.
Stephen Kuhn and Mary Stiner of the University of Arizona suggest an
alternative hypothesis. They believe that modern humans entered Europe
with cultures having a division of labor that was less risky for pregnant
women, mothers, and young children. They think that women mostly
collected vegetables, fruits, and nuts, while men concentrated on the far more
hazardous task of hunting large animals. The occasional loss of
men in hunting accidents would not have significantly affected birth rates
and the survival of children. As a consequence, there would have been
a steady growth in the size of modern human populations. In contrast,
Kuhn and Stiner believe that both men and women among the Neandertals were
involved in the dangerous hunting of big game animals with weapons that
required close encounters with their prey. As a result, their
populations were kept low and relatively uncompetitive.

Reconstruction of
a
late Neandertal child's
appearance based
on skeletal material
found in Gibraltar

Another
provocative hypothesis to explain the disappearance of Neandertals has been
proposed by Leslie Aiello. It is connected with the fact that
Neandertals progressively became scarcer as Europe moved into the coldest
phase of the last ice age. She suggests that Neandertal habitation
sites were limited to areas in which winter temperatures did not go below 0º F.
(-18º C.) because their technology was not up to
dealing with harsher conditions. These relatively warmer areas would
have become more and more isolated valley pockets as the climate cooled,
especially after 30,000
years ago. Eventually, they
would have disappeared and the Neandertals within them would have perished
from the cold. Aiello believes that the modern humans living in Europe at
that time survived because their technology for dealing with extreme ice age
conditions was superior. This hypothesis provides a logical explanation
for the disappearance of most but not all
Neandertals. It does not explain why the Neandertals living in the
comparatively warmer areas of Southwest Asia also became extinct. It also
is based on the questionable assumption that Neandertals were not smart or
inventive enough to develop adequate technology for severe cold winter
conditions. However, it is true that around 35,000
years ago the known
Neandertal habitation sites began to disappear from Central Europe.
After 30,000 years ago, they apparently only survived in what would have
been the relatively warmer southern European regions of Portugal, Spain, and
Gibraltar, and they ended there by about 28,000-24,000 years ago.
There was a major climatic shift to much colder conditions around 24,000
years ago.

Regardless of how they are classified
or what happened to them, it has now
become clear that the Neandertals were a very successful
line of humans for 300-400,000 years. They were the first
people to live successfully
in subarctic environments of the northern
hemisphere during
at least the milder phases of an ice age.
They first appeared in most of Europe during an interglacial when the climate was
similar to today. With the onset of the last ice age about 75,000 years
ago, some Neandertals may have migrated to Southwest Asia (at least to Israel and Iraq)
where it was warmer. However, others
adapted to the increasingly severe climatic conditions of Europe. They did this
primarily with new cultural inventions discussed in the next section
of this tutorial.

Other
Archaic Human Populations

Mitochondrial
DNA nucleotide sequence differences between
Neandertals indicate that very likely there were three different geographical populations of
them. These sub-groups lived in Western Europe, Southern Europe, and
Southwestern Asia. It also is becoming clear that the Neandertals were not
the only archaic human population in Eurasia. Mitochondrial
DNA recovered from a 50,000-30,000 year old finger bone found at a Siberian
cave site known as Denisova is from
an up to now unknown form of human (referred to as the Denisovans ).
This possible new variety or even new species of archaic human lived at the
same time as Neandertals and early modern humans. We now know that the
Denisovans interbred with the early Homo sapiens whose descendents
would become South Asians and Southwest Pacific Islanders. About 1% of the DNA of some populations in South
China today and 4-6% of the DNA of
New Guineans and other Melanesians was inherited from the Denisovans.

NEWS:
Clive Finlayson of the Gibraltar Museum reported in the
September 6, 2006 issue of Naturethat there was a Neandertal occupation of Gorham's Cave in Gibraltar from
32,000 to 24,000 years ago. He believes that Neandertals and
modern humans lived simultaneously in Gibraltar and nearby areas of Southern
Spain for several thousand years. If his radiocarbon dates are
correct, it means that Neandertals persisted in Southern Europe for upwards
of 4,000 years after they are generally thought to have disappeared.